Keywords:

Prickly pear

Cladode

Propagation

Forage

Eect of sectioning, drying, and shading on the propagation of Opuntia cochenillifera (L.) Mill.

cladodes

Efecto del seccionamiento, secado y sombreado en la multiplicación de cladodios de Opuntia

cochenillifera (L.) Mill.

Efeito da divisão, secagem e sombreamento na multiplicação de cladódios de Opuntia cochenillifera

(L.) Mill.

Rafael Andrés Llinás Vergara

Carlos Alberto Atencio León

Andy José González Atencio

Gisela Rivero Maldonado

Jorge Ortega Alcalá

Ciolys Colmenares

Ruben León

Rev. Fac. Agron. (LUZ). 2026, 43(2): e264325

ISSN 2477-9407

DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v43.n2.VII

Crop production

Associate editor: Dr. Jorge Vilchez-Perozo

University of Zulia, Faculty of Agronomy

Bolivarian Republic of Venezuela

Facultad de Agronomía, Universidad del Zulia, Venezuela.

Departamento de Botánica, Facultad de Agronomía,

Universidad del Zulia, Venezuela.

Departamento de Estadística, Facultad de Agronomía,

Universidad del Zulia, Venezuela.

Received: 07-02-2026

Accepted: 26-03-2026

Published: 28-04-2026

Abstract

Opuntia cochenillifera (L.) Mill. has a forage potential in

arid areas. Because pastures are insucient to meet livestock’s

nutritional demands during the dry season, this research aimed

to evaluate the eects of sectioning (S), drying (D), and shading

(Sh) of cladodes on the propagation of Opuntia cochenillifera (L.)

Mill. The S factors (whole cladode and apical, middle, and basal

sections), D (pre-drying at room temperature on mesh under shade

for 14 days), and Sh (with and without shading under 80 % density

raa mesh), resulted in a combination of 16 treatments, arranged

in a split-plot treatment design and a completely randomized

experimental design. Percentages of living, rooted, and sprouted

cladodes, number of shoots per cladode, and number and length

of roots were evaluated 35 days after establishment. An analysis

of variance and a means test were applied to determine dierences

among treatments. An increase in the percentage of rooted cladodes

was obtained with the middle, apical, and basal fractions compared

to the whole cladode, regardless of the drying process. Prior shading

was counterproductive for cladode survival; high environmental

humidity, combined with their water content, may have favored the

appearance of secondary rots. In general, considering most of the

variables evaluated, cladode sectioning without drying emerged as

the best technique for asexual propagation of the species, due to the

yield of vegetative material and its practicality.

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

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2-6 |

Resumen

La Opuntia cochenillifera (L.) Mill. tiene potencial forrajero en

zonas áridas. Debido a que en la época seca los pastizales no son

sucientes para satisfacer la demanda alimenticia del ganado, en

esta investigación se planteó el objetivo de evaluar el efecto del

seccionamiento (S), secado (Se) y sombreado (So) de los cladodios

en la multiplicación de Opuntia cochenillifera (L.) Mill. Los factores

S (cladodio entero y secciones apical, media y basal), Se (secado

previo a temperatura ambiente sobre malla y bajo sombra durante 14

días) y So (con y sin sombreado bajo malla raa al 80 %) resultaron

en una combinación de 16 tratamientos, dispuestos en un diseño de

tratamiento de parcelas divididas y un diseño experimental totalmente

al azar. Se evaluaron porcentajes de cladodios vivos, enraizados y

brotados, número de brotes por cladodio y número y longitud de raíces

a los 35 días del establecimiento. Se utilizó un análisis de varianza y

prueba de medias para determinar diferencias entre tratamientos. Se

obtuvo un incremento en el porcentaje de cladodios enraizados con

la fracción medial, fracción apical y fracción basal, en comparación

con el cladodio entero, independientemente del secado. El sombreado

previo fue contraproducente para la sobrevivencia de los cladodios; la

alta humedad ambiental, aunado al contenido hídrico de los mismos

pudo favorecer la aparición de pudriciones secundarias. En general,

considerando la mayoría de las variables evaluadas el seccionamiento

del cladodio, sin secado se perló como la mejor técnica para

propagar asexualmente la especie, debido al rendimiento del material

vegetativo, así como por su practicidad.

Palabras clave: nopal, cladodio, propagación, forraje.

Resumo

A Opuntia cochenillifera (L.) Mill tem potencial como forragem

em áreas áridas. Devido ao décit de sementes, foi testado um método

de propagação assexuada através dos seguintes fatores de estudo:

sombreamento (com e sem sombreamento com tela de ráa de 80 %),

seccionamento (cladódio inteiro e seções apical, média e basal) e pré-

secagem (à temperatura ambiente sobre tela e à sombra por 14 dias),

resultando em uma combinação de 16 tratamentos, dispostos em um

delineamento experimental de parcelas divididas e blocos aleatórios.

As porcentagens de cladódios vivos, enraizados e brotados, o número

de brotos por cladódios e o número e comprimento das raízes foram

avaliados 35 dias após o estabelecimento. Análise de variância e

testes de média foram aplicados para determinar as diferenças entre

os tratamentos. Um aumento na porcentagem de cladódios enraizados

foi obtido com a fração medial, fração apical e fração basal, em

comparação com o cladódio inteiro, independentemente da secagem.

O sombreamento prévio foi contraproducente para a sobrevivência

dos cladódios; a alta umidade ambiental, combinada com o teor de

água dos cladódios, pode ter favorecido o aparecimento de podridão

secundária. Em geral, considerando as variáveis avaliadas, o corte

dos cladódios sem secagem emergiu como a melhor técnica para a

propagação assexuada da espécie, devido ao rendimento do material

vegetativo e sua praticidade.

Palavras-chave: go-da-Índia, cladódio, propagação, forragem.

Introduction

Livestock remains the main source of income for rural

communities in arid zones and is a critical component of resilient

production systems and an indicator of wealth. However, this sector

faces numerous challenges, including food availability and climate

change. Grasslands in semi-arid regions are vital for livestock

production systems, although their contribution to animal feed is

declining. This situation has increased interest in the use of CAM

species as forage due to their high eciency in the use of water

and ability to be cultivated on abandoned lands with marginal soils

(Niechayev et al., 2019). In addition, CAM species oer additional

benets, as they are ideal for reforestation and ecological restoration

in areas aected by desertication and climate change (Owen et

al., 2015). In this context, the prickly pear cactus (Opuntia spp.)

emerges as a promising solution; it is not only drought-resistant

but also converts biomass more eciently than C3 and C4 plants.

In fact, it generates three times more biomass per unit of water than

C4 plants and ve times more than C3 plants. It is estimated that

900,000 hectares of prickly pear are cultivated for forage worldwide

(Reynolds and Arias Jiménez, 2003). In Venezuela, the prickly pear

of the genus Opuntia is one of the cacti with the greatest presence in

semi-arid regions (Gallardo et al., 2016). It has been observed that

the species Opuntia cochenillifera (L.) Mill., which lacks spines,

has a high preference and palatability by cattle; it stands out for its

high nutritional value, with a crude protein content of 6.2 % and

a digestibility of 78 %. Its adaptability to various environmental

conditions makes it a sustainable option for livestock production

systems, especially in arid and semi-arid regions (Vázquez Mendoza

et al., 2019). However, the problem lies in the insucient availability

of propagules for their massive propagation. This underscores the

design of strategies to develop livestock production systems based on

prickly pear, favoring the human population and reducing pressure on

natural grasslands. In this sense, propagation techniques are essential

for the mass propagation and establishment of plantations of this

species.

It has been established that reducing the size of the cutting of the

cladode does not reduce its potential for rhizogenesis and caulogenesis.

This fragmentation represents a substantial gain in material and time,

especially for large, cultivated areas of O. cus-indica (Stambouli-

Essassi et al., 2015). While Mondragón et al., (2003) suggest that the

size of the cladode does not aect its ability to form roots and shoots,

Homrani et al., (2016) indicate a positive correlation with its number.

Another factor evaluated in asexual or vegetative propagation is the

pre-drying of cladodes, as it reduces water loss and the incidence

of rot, favoring rooting (Mulas and Dessena, 2019). Therefore, the

objective of this study was to evaluate the eect of the factors of

pre-drying, sectioning, and shading of the cladodes, as well as their

interactions, on the propagation of Opuntia cochenillifera (L.) Mill.,

to provide an alternative forage source in arid and semi-arid regions.

Materials and methods

Experiment location

The trial was carried out at the “El Mamonal” farm, located on

the Falcón-Zulia highway, Miranda municipality, Zulia state, Piñero

sector, coordinates 10º39´57.3”N and 71º21´47.2”W, belonging to

a Tropical Dry Forest ecosystem, with an average temperature of

28.7°C and average annual rainfall of 300 mm. Rainfall is distributed

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Llinás et al. Rev. Fac. Agron. (LUZ). 2026, 43(2): e264325

3-6 |

irregularly, presenting a bimodal behavior. This means that there are

two rainfall peaks, one in May and one in October, while the months

of December to March and July to August are the driest periods

(Ferrer-Paris et al., 2015).

Plant material

Cladodes of Opuntia cochenillifera (L.) Mill. (Cactaceae) were

used, selected from plants approximately 1 to 2 years old and in good

phytosanitary condition, from a population of 800 prickly pear cacti

established in cultivation at the production facility. The cladodes

measured approximately 28 cm in length and 11 cm in diameter.

Planting substrate

The cladodes were planted in 20 kg bags with a substrate mixture

composed of two parts topsoil and one part of previously washed

cow manure (2:1 ratio). In each bag, the cladodes were positioned

vertically, respecting their natural polarity, and buried to a depth of

up to two-thirds (2/3) of their length (Figure 1).

Figure 1. Establishment of the cladodes in bags.

Cultural practices

The cladodes were irrigated twice a week, in the afternoon,

with a total contribution of 400 cc per bag. The irrigation frequency

was every three days, adjusted according to atmospheric conditions

to ensure that the substrate was maintained at eld capacity. Weed

control was performed manually.

Study factors and levels

Three factors at dierent levels were studied: Sectioning of the

cladodes: whole cladode and apical, middle, and basal sections; each

section measured approximately 9 cm. The sections were made with

sharp blades, disinfecting them at the beginning with 0.5 % sodium

hypochlorite for 10 minutes. There was no application of healing

agents in the cut area, considering the good healing observed in

previous trials.

Drying of the cladodes: without drying and with drying.

Both whole and sectioned cladodes were left to dry at room

temperature on mesh and under shade for 14 days.

Shading of the cladodes: without shade and with shade.

The cladodes established in shade had a roofed area covered with

80 % density raa mesh. The cladodes without shade were placed

in the eld under full sun exposure. In both cases, two rows of eight

bags were placed randomly within each row.

Treatments

From the combination of the levels of study, 16 treatments were

generated. The experimental unit consisted of a bag where four

cladodes (whole, apical, middle, and basal) were established, with

two replications per treatment, for a total of 32 experimental units.

Experimental design

The treatment design used was Split Plots 2x4x2, whose

combinations were established in a completely randomized

experimental design. The shading factor was located in the main plot,

and the sectioning and drying factors were in the secondary plots.

Statistical processing and analysis

An analysis of variance was performed, and for the eects that

were signicant, Tukey’s means test was applied, using the InfoStat

software under the Windows platform.

Results and discussion

The following results were obtained based on the eects of

sectioning and drying the cladodes under full sun exposure, since the

cladodes placed in the shade were not viable, resulting in one hundred

percent mortality.

Drying under shade was carried out from February 1st to 15th, 2025

(14 days), where a relative humidity of 18 % corresponding to the dry

season prevailed; however, there was a generalized rot in the Opuntia

cladodes, possibly associated with the environmental conditions during

shading (under Polyshade mesh) and exacerbated by the weekly 2-mm

irrigation depth, prevailing a microclimate of high humidity and limited

ventilation, which caused the proliferation of fungal and bacterial

pathogens.

It has been reported that shade has a positive inuence on the

growth and biomass production of the prickly pear cactus. Compared

to full sun conditions, shading up to 50 % benets plant establishment,

leaf area, and biomass production. However, higher shading (75 %)

has a negative eect on plant survival due to microbial rot and a lower

reduction in dry biomass (Dev et al., 2018). This suggests that shading

levels should be measured with greater precision in future trials.

Percentage of rooted cladodes (PRC)

The analysis of variance showed signicant dierences (P<0.05)

only for the eect of sectioning on the PRC variable. Drying as a single

eect and its interaction with sectioning were not signicant (P>0.05).

Regarding sectioning, the highest PRC was achieved in the middle

and apical sections, both reaching 100 % rooted cladodes. These did

not dier statistically from the basal section (93.51 %) but did show

signicant dierences compared to the whole cladodes (62.50 %),

which exhibited the lowest PRC, as shown in Figure 2.

Figure 2. Eect of sectioning on the variable percentage of

cladodes rooted in Opuntia cochenillifera (L.) Mill.

MS: Middle section; AS: Apical section; BS: Basal

section; WC: Whole cladode. Equal letters do not present

statistically signicant dierences (P>0.05).

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Rev. Fac. Agron. (LUZ). 2026, 43(2): e264325 April-June ISSN 2477-9409.

4-6 |

average number of roots per cladode did not vary signicantly among

the dierent drying treatments.

The values recorded for the number of roots per cutting ranged

from 1.87 for the 96-hour drying treatment to 2.01 for the 120-hour

drying treatment. Although the proportion of rooted cuttings was

higher with 96 hours of drying (92.6 %) compared to 72 hours (85.7

%) and 120 hours (83.9 %), the number of individual roots per cutting

did not show a direct dependence on these drying times.

Root formation in Opuntia species, such as prickly pear (Opuntia

cus-indica) and O. robusta, is markedly inuenced by the planting

method. According to Snyman (2006), when the cladode was placed

at on the ground, both species developed a greater number of

roots. This phenomenon is due to an increase in the contact of the

areoles, which are the structures from which the roots sprout, with

the substrate. Variations in the shape and size of cladodes between O.

cus-indica and O. robusta explain the dierences in the number of

areoles that come into contact with the soil and, consequently, in root

development. Despite these morphological dierences, most of the

areoles in contact with the ground in both species managed to form

roots after three weeks, regardless of the planting method used.

Root length (RL)

For RL per cladode, the analysis of variance detected signicant

dierences (P<0.05) due to the eect of sectioning and drying. The

interaction of both factors was not signicant (P>0.05).

In reference to sectioning, the middle fraction (7.63 cm) diered

from that of the whole cladode (3.98 cm). The apical and basal

fractions had a similar behavior to the other fractions, as shown in

Figure 4.

Figure 4. Eect of sectioning on the variable root length in

Opuntia cochenillifera (L.) Mill. MS: Cladode in middle

section; AS: Cladode in apical section; BS: Cladode in basal

section; WC: Whole cladode. Equal letters do not present

statistically signicant dierences (P>0.05)

Similarly, signicant dierences (P<0.05) were observed due to

the eect of drying, as shown in Figure 5. The cladodes that were

not dried obtained a mean of 6.80 cm, while those subjected to the

procedure reached 4.53 cm, a 66.61% lower response (P<0.05). This

fact strengthens the results obtained for the percentage of cladodes

rooted, which suggests that drying under shade does not have a

determining eect on root production.

The above is explained by the cells’ need for water to support

their metabolic processes, such as sustaining cell multiplication and

elongation, leading to the production of tissues responsible for root

growth (Khanna, 2024).

According to Stambouli-Essassi et al., (2015), sectioning of

cladodes, even in small portions such as halves, quarters, and tenths,

has proven to be an eective strategy for vegetative propagation of

Opuntia cus-indica. This technique does not reduce the potential of

cladodes for rhizogenesis. According to these authors, sectioning can

lead to 100 % rooting in the cuttings of right cladodes, with a higher

number of roots (80) developing in specic portions such as the right

basal tenth. The highest percentage of secondary cladodes (70 % and

74 %) was initiated in the right apical cuttings, and the longest roots

(13 and 14 cm) were initiated in the two basal tenths, both left and

right. This method represents a considerable advantage in terms of

material and time optimization, which is especially benecial for the

establishment of large prickly pear plantations.

Likewise, Mulas and Dessena (2019) evaluated the feasibility

of the propagation of Opuntia cus-indica using cladode fragments;

they concluded that the eect of sectioning combined with a pre-

drying process is a benecial methodology for the propagation of

the prickly pear cactus. In contrast to the results obtained in this

research, Mulas and Dessena (2019) showed that cladode fragments

that were subjected to 96 hours of drying not only managed to root,

but crucially, showed an increase in their initial fresh weight after

rooting. This weight increase is a key indicator of successful and

vigorous plant establishment, suggesting that this approach allows

for a higher number of viable and well-developed individuals to be

obtained from the same amount of mother material, thus optimizing

the rate of plant propagation.

Number of roots per cladode (NRC)

The analysis of variance did not detect signicant dierences

(P>0.05) for any of the eects evaluated (drying and sectioning), nor for

their interaction. The overall average was 8.94 ± 4.07 roots.cladode

-1

Figure 3 shows a rooted cladode, showing the approximate number

of roots obtained.

Figure 3. Cladode with root formation

The previous results are comparable to those obtained by

Mulas and Dessena (2019), who, while evaluating the propagation

of Opuntia cus-indica from cladode fragments, observed that the

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Llinás et al. Rev. Fac. Agron. (LUZ). 2026, 43(2): e264325

5-6 |

corresponding to the basal and middle sections, which obtained 1.63

and 1.06 shoots.cladode

-1

, respectively, in the NSC of the apical

section, a similar behavior is observed to the rest of the levels studied,

whose level produced 1.94 shoots.cladode

-1

(Figure 7). Figure 8

shows a whole cladode with cauline shoots.

Figure 5. Eect of drying on the variable root length in Opuntia

cochenillifera (L.) Mill. DC: Dry cladodes; CWD:

Cladodes without drying. Equal letters do not present

statistically signicant dierences (P>0.05).

The sectioning of Opuntia cus-indica cladodes has a signicant

inuence on the root length. According to the observations of

Stambouli-Essassi et al., (2015), whole cladodes transplanted in

spring produce the longest roots, reaching 17 cm, while cladode

halves under the same conditions develop 13 cm roots. In the case of

the cladodes sectioned into quarters, the right portions, both basal and

apical portions, generated longer roots (9 cm and 8.8 cm, respectively)

compared to the left portions (6.8 cm and 7.8 cm). The basal tenths of

the cladode, especially those on the right side, produced the longest

roots, reaching 14 cm, while the middle portions showed the shortest

roots (between 5 and 6 cm). These ndings suggest that reducing

the size of the cladode does not necessarily decrease its rhizogenesis

potential, and that planting polarity inuences root development.

Stambouli-Essassi et al., (2015) also concluded that right-side

cladode portions, planted vertically and with their normal polarity,

demonstrated the best results in aptitude for rhizogenesis and

secondary cladode initiation, compared to left-side portions planted

horizontally.

Percentage of sprouted cladodes (PSC)

Like the PRC variable, the analysis of variance showed only

signicant eects (P<0.05) for the sectioning factor on the PSC

variable. In contrast, drying and the interaction of drying and

sectioning were not signicant (P>0.05).

The highest PSC was obtained by the whole cladode (100 %),

being statistically dierent (P<0.05) from the middle section (62.50 %),

and the lowest PSC value. The basal (87.50 %) and apical (81.25 %)

sections had a similar behavior among themselves and with the rest of

the levels studied (Figure 6).

These results are similar to those obtained by Stambouli-Essassi

et al., (2015) who determined dierences in sprouting percentages

according to the type of sectioning; the whole cladodes reached 100

% sprouting, while those sectioned in two halves showed 90 % for

the apical segments and 60 % sprouting for the basal segments; the

cladodes sectioned into four parts presented the lowest percentages

(20-30 %). These ndings suggested that excessive sectioning of the

cladode compromises sprouting capacity, possibly due to the reduction

of nutrient reserves and available water, as well as an increased risk of

dehydration and susceptibility to pathogens.

Number of shoots per cladode (NSC)

The analysis of variance detected signicant dierences (P<0.01)

due to the eect of the sectioning factor on the variable NSC.

The highest number of shoots corresponded to the whole cladode

with 3.13 shoots.cladode

-1

, which diered (P<0.05) from those

Figure 6. Eect of sectioning on the variable percentage of cladodes

sprouted in Opuntia cochenillifera (L.) Mill. WC: Whole

cladode; BS: Cladode in basal section; AS: Cladode in

apical section; MS: Cladode in middle section; Equal letters

do not present statistically signicant dierences (P>0.05).

Figure 7. Eect of sectioning on the variable number of shoots per

cladode in Opuntia cochenillifera (L.) Mill. WC: Whole

cladode; BS: Cladode in basal section; AS: Cladode in

apical section; MS: Cladode in middle section; Equal letters

do not present statistically signicant dierences (P>0.05).

In the research conducted by Mateus-Arango (2018), the

sectioning of the cladodes of Opuntia cus-indica proved to be an

eective strategy to increase the average number of shoots; thus,

complete cladodes generated an average of 1.83 shoots; those divided

into two parts produced 3.25 shoots, and those sectioned into four

parts reached an average of 4.75 shoots. This upward trend suggested

that cladode fragmentation enhances the activation of a greater

number of latent buds, resulting in higher proliferation of shoots per

sectioned unit. The results of Mateus-Arango (2018) are dierent

from those obtained in this research, where the fragmentation of the

cladodes caused a decrease in the sprouting percentage.

Khalafalla et al., (2007) evaluating the in vitro micropropagation

of Opuntia cus-indica, highlighted the high shoot multiplication rate

obtained, reporting a generation of 26.5 ± 1.74 shoots per explant

in a period of 90 days, using an MS medium enriched with 5 mg.L

-1

benzyladenine. This result underlines the potential of tissue culture

techniques for the mass production of plant material, as they allow for

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Rev. Fac. Agron. (LUZ). 2026, 43(2): e264325 April-June ISSN 2477-9409.

6-6 |

obtaining a large number of shoots from an initial fragment. In relation

to the previous results, it is convenient to point out that this eciency

was achieved in a highly controlled laboratory environment, inherent

to micropropagation, diering from the conventional propagation

conditions under which this research was carried out; however, it

could represent an excellent alternative for the propagation of the

evaluated species.

Conclusions

Cladode sectioning proved to be the most eective technique for

rooting and thus for the mass propagation of O. cochenillifera (L.)

Mill. However, the use of whole cladodes allows for an increase in

the number of shoots, which is a factor to consider depending on the

objectives of propagation.

Pre-drying had an impact on the obtaining of cladodes with longer

root lengths, which can be benecial for their survival; however, as

it did not have a signicant impact on the other variables studied,

its application should continue to be evaluated, since it implies an

additional expenditure of time and resources.

Shading using 80 % density raa mesh proved to be

counterproductive due to the high incidence of pathogens; therefore,

reducing this percentage is suggested to allow for greater direct solar

radiation. Consequently, the implementation of meshes with lower

shade density (30 – 50 %) is recommended, as well as evaluating other

aeration and temperature conditions.

With the results obtained, a mass propagation method of O.

cochenillifera (L.) Mill. was established to ensure forage availability

during dry seasons, providing a foundation for the sustainability of

livestock production systems in arid and semi-arid environments.

Acknowledgment

The authors wish to extend their sincere gratitude to the “El

Mamonal” farm and to Dr. Eleazar Soto Belloso for providing their

facilities and the Opuntia mother plant stock, as well as the supplies and

materials necessary to conduct the experimental phase of this research.

Literature cited

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and biomass production of cactus (Opuntia cus-indica (L.) Mill.).

International Journal of Current Microbiology and Applied Sciences

(IJCMAS), 7, 3145–3153. https://www.ijcmas.com/special/7/Rahul%20

Dev,%20et%20al.pdf

Ferrer-Paris J.R., I. Stachowicz, L. Morán, L. Gonz´alez, AY Sánchez-Mercado,

C. Lozano, J. Soto, A. Cardozo-Urdaneta y T. (2015). Caracterización

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Figure 8. Whole cladode with cauline shoots.